''Chenopodium'' is a

genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...

of numerous species of

perennial In horticulture, the term perennial ('' per-'' + '' -ennial'', "through the year") is used to differentiate a plant from shorter-lived annuals and biennials. It has thus been defined as a plant that lives more than 2 years. The term is also ...

or annual herbaceous

flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Ancient Greek, Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed with ...

s known as the goosefoot, which occur almost anywhere in the world. It is placed in the family

Amaranthaceae Amaranthaceae ( ) is a family of flowering plants commonly known as the amaranth family, in reference to its type genus '' Amaranthus''. It includes the former goosefoot family Chenopodiaceae and contains about 165 genera and 2,040 species, maki ...

in the

APG II system The APG II system (Angiosperm Phylogeny Group II system) of plant classification is the second, now obsolete, version of a modern, mostly Molecular phylogenetics, molecular-based, list of systems of plant taxonomy, system of plant taxonomy that ...

; older classification systems, notably the widely used

Cronquist system The Cronquist system is a list of systems of plant taxonomy, taxonomic classification system of angiosperms, flowering plants. It was developed by Arthur Cronquist in a series of monographs and texts, including ''The Evolution and Classification of ...

, separate it and its relatives as

Chenopodiaceae Amaranthaceae ( ) is a family of flowering plants commonly known as the amaranth family, in reference to its type (biology), type genus ''Amaranthus''. It includes the former goosefoot family Chenopodiaceae and contains about 165 genera and 2,040 ...

, but this leaves the rest of the Amaranthaceae

polyphyletic A polyphyletic group is an assemblage that includes organisms with mixed evolutionary origin but does not include their most recent common ancestor. The term is often applied to groups that share similar features known as Homoplasy, homoplasies ...

. However, among the Amaranthaceae, the genus ''Chenopodium'' is the namesake member of the subfamily

Chenopodioideae The Chenopodioideae are a subfamily of the flowering plant family Amaranthaceae in the APG III system, which is largely based on molecular phylogeny, but were included – together with other subfamilies – in the family Chenopodiaceae, or goose ...

Description

The species of ''Chenopodium'' (s.str., description according to Fuentes et al. 2012) are annual or

herbs,

shrubs A shrub or bush is a small to medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from trees by their multiple ...

or small

trees In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, e.g., including only woody plants with secondary growth, only p ...

. They generally rely on

alkaline soil Alkali, or alkaline, soils are clay soils with high Soil pH, pH (greater than 8.5), a poor soil structure and a low infiltration capacity. Often they have a hard calcareous layer at 0.5 to 1 metre depth. Alkali soils owe their unfavorable ...

. They are nonaromatic, but sometimes fetid (foul-smelling). The young stems and leaves are often densely covered by vesicular globose hairs, thus looking farinose. Characteristically, these

trichomes Trichomes (; ) are fine outgrowths or appendages on plants, algae, lichens, and certain protists. They are of diverse structure and function. Examples are hairs, glandular hairs, scales, and papillae. A covering of any kind of hair on a plant ...

persist, collapsing later and becoming cup-shaped. The branched stems grow erect, ascending, prostrate or scrambling. Lateral branches are alternate (the lowermost ones can be nearly opposite). The alternate or opposite

leaves A leaf (: leaves) is a principal appendage of the stem of a vascular plant, usually borne laterally above ground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, stem, ...

are petiolate. Their thin or slightly fleshy leaf blade is linear, rhombic or triangular- hastate, with entire or dentate or

lobed The following terms are used to describe leaf plant morphology, morphology in the description and taxonomy (biology), taxonomy of plants. Leaves may be simple (that is, the leaf blade or 'lamina' is undivided) or compound (that is, the leaf blade ...

margins.

Inflorescence In botany, an inflorescence is a group or cluster of flowers arranged on a plant's Plant stem, stem that is composed of a main branch or a system of branches. An inflorescence is categorized on the basis of the arrangement of flowers on a mai ...

s are standing terminal and

lateral Lateral is a geometric term of location which may also refer to: Biology and healthcare * Lateral (anatomy), a term of location meaning "towards the side" * Lateral cricoarytenoid muscle, an intrinsic muscle of the larynx * Lateral release ( ...

. They consist of spicately or paniculately arranged glomerules of flowers. Plants are

monoecious Monoecy (; adj. monoecious ) is a sexual system in seed plants where separate male and female cones or flowers are present on the same plant. It is a monomorphic sexual system comparable with gynomonoecy, andromonoecy and trimonoecy, and contras ...

(rarely

dioecious Dioecy ( ; ; adj. dioecious, ) is a characteristic of certain species that have distinct unisexual individuals, each producing either male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproduction is ...

). In monoecious plants flowers are dimorphic or pistillate. Flowers consist of (4–) 5

perianth The perianth (perigonium, perigon or perigone in monocots) is the non-reproductive part of the flower. It is a structure that forms an envelope surrounding the sexual organs, consisting of the calyx (sepals) and the corolla (petals) or tepal ...

segments connate, basally (at the base) or close to the middle, usually membranous margined and with a roundish to keeled back; almost always 5 stamens, and one ovary with 2 stigmas. In fruit, perianth segments become sometimes coloured, but mostly keep unchanged, somewhat closing over or spreading from the fruit. The

pericarp Fruits are the mature ovary or ovaries of one or more flowers. They are found in three main anatomical categories: aggregate fruits, multiple fruits, and simple fruits. Fruitlike structures may develop directly from the seed itself rather th ...

is membranous or sometimes succulent, adherent to or loosely covering the seed. The horizontally oriented seeds are depressed-globular to lenticular, with rounded to subacute margin. The black seed coat is almost smooth to finely striate, rugulose or pitted.

Taxonomy

The genus ''Chenopodium'' was described by

Carl Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...

in 1753 (In: ''Species Plantarum'', Vol. 1, p. 218–222). Type species is '' Chenopodium album''. This generic name is derived from the particular shape of the leaf, which is similar to a goose's foot: from

Greek Greek may refer to: Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group *Greek language, a branch of the Indo-European language family **Proto-Greek language, the assumed last common ancestor of all kno ...

χήν (''chen''), "goose" and πούς (''pous''), "foot" or (''podion''), "little foot". In its traditional circumscription, ''Chenopodium'' comprised about 170 species. Phylogenetic research revealed, that the genus was highly

and did not reflect how species were naturally related. Therefore, a new classification was necessary. Mosyakin & Clemants (2002, 2008) separated the glandular species as genus '' Dysphania'' (which includes

epazote ''Dysphania ambrosioides'', formerly ''Chenopodium ambrosioides'', known as epazote, Jesuit's tea, Mexican tea or wormseed, is an annual or short-lived perennial herb native to the Americas. Description ''Dysphania ambrosioides'' is an annual p ...

) and '' Teloxys'' in tribe Dysphanieae. Fuentes-Bazan et al. (2012) separated many species to genera ''

Blitum ''Blitum'' is a genus of flowering plants in the amaranth family Amaranthaceae, subfamily Chenopodioideae. It is closely related to genus '' Spinacia''. Its 12 species were traditionally placed in the genera '' Chenopodium'', ''Monolepis'', or '' ...

'' (in tribe Anserineae), '' Chenopodiastrum'', '' Lipandra'', and '' Oxybasis'' (like ''Chenopodium'' in tribe Atripliceae). They included ''Rhagodia'' and ''Einadia'' in ''Chenopodium''.

Species

132 species are currently accepted. * '' Chenopodium acerifolium'' * '' Chenopodium aciculare'' * '' Chenopodium acuminatum'' Willd. * '' Chenopodium adpressifolium'' * '' Chenopodium albescens'' * '' Chenopodium album'' – white goosefoot, nickel greens, dungweed, ''bathua'', ''chandali'', ''chandaliya'', fat hen, lamb's quarters, pigweed ** ''Chenopodium album'' var. ''album'' (synonym '' Chenopodium strictum'' Roth) * '' Chenopodium allanii'' * '' Chenopodium atripliciforme'' * '' Chenopodium atrovirens'' – dark goosefoot, pinyon goosefoot * '' Chenopodium attenuatum'' * '' Chenopodium aureum'' – golden goosefoot * '' Chenopodium auricomiforme'' * '' Chenopodium auricomum'' – Queensland bluebush * '' Chenopodium ayare'' * '' Chenopodium baccatum'' (Syn. ''Rhagodia baccata'') * '' Chenopodium benthamii'' (Syn.: ''Rhagodia latifolia'') * ''

Chenopodium berlandieri ''Chenopodium berlandieri'', also known by the common names pitseed goosefoot, lamb's quarters (or lambsquarters), and ''huauzontle'' (Nahuatl) is an annual plant, annual herbaceous plant in the family Amaranthaceae. The species is widespread i ...

Chenopodium neomexicanum ''Chenopodium'' is a genus of numerous species of perennial or annual plant, annual herbaceous flowering plants known as the goosefoot, which occur almost anywhere in the world. It is placed in the family Amaranthaceae in the APG II system; old ...

'' – New Mexico goosefoot * '' Chenopodium nesodendron'' * '' Chenopodium nevadense'' – Nevada goosefoot * '' Chenopodium nitens'' * '' Chenopodium nitrariaceum'' (F.Muell.) F.Muell. ex Benth. – nitre goosefoot * '' Chenopodium nitens'' – shiny goosefoot * '' Chenopodium novopokrovskyanum'' * '' Chenopodium nutans'' (Syn.: ''Rhagodia nutans'') * '' Chenopodium oahuense'' – ''Āheahea'' (

Hawaii Hawaii ( ; ) is an island U.S. state, state of the United States, in the Pacific Ocean about southwest of the U.S. mainland. One of the two Non-contiguous United States, non-contiguous U.S. states (along with Alaska), it is the only sta ...

) * '' Chenopodium obscurum'' * '' Chenopodium opulifolium'' Schrad. ex W.D.J.Koch & Ziz – grey goosefoot * '' Chenopodium pallescens'' – pallid goosefoot * ''

Chenopodium pallidicaule ''Chenopodium pallidicaule'', known as cañihua, canihua or cañahua (from Quechua language, Quechua 'qañiwa, qañawa or qañawi') and also kañiwa or kaniwa, is a species of goosefoot, similar in character and uses to the closely related quinoa ...

Chenopodium stenophyllum ''Chenopodium'' is a genus of numerous species of perennial or annual herbaceous flowering plants known as the goosefoot, which occur almost anywhere in the world. It is placed in the family Amaranthaceae in the APG II system; older classifica ...

'' * '' Chenopodium striatiforme'' * '' Chenopodium subficifolium'' * '' Chenopodium subglabrum'' – smooth arid goosefoot, smooth goosefoot * '' Chenopodium suecicum'' – green goosefoot * '' Chenopodium × thellungii'' * '' Chenopodium tonkinense'' * '' Chenopodium triandrum'' (Syn.: ''Rhagodia triandra'') * '' Chenopodium × tridentinum'' * '' Chenopodium × trigonocarpum'' * '' Chenopodium trigonon'' (Syn.: ''Einadia trigonos'') * '' Chenopodium twisselmannii'' – Twisselmann's goosefoot, high meadow goosefoot * '' Chenopodium ulbrichii'' * '' Chenopodium ulicinum'' * '' Chenopodium × unarii'' * '' Chenopodium × variabile'' (''C. album × C. berlandieri'') * '' Chenopodium vulvaria'' – stinking goosefoot, notchweed * '' Chenopodium wahlii'' – Wahl's goosefoot * '' Chenopodium watsonii'' – Watson's goosefoot * '' Chenopodium wilsonii'' (Syn.: ''Rhagodia crassifolia'') * '' Chenopodium × zahnii'' * '' Chenopodium zerovii'' * '' Chenopodium zoellneri''

Excluded species

* ''

'' (12 species): ** '' Blitum bonus-henricus'' – Good King Henry, perennial goosefoot, poor-man's asparagus, Lincolnshire spinach, markery ** '' Blitum californicum'' – California goosefoot, Indian lettuce ** '' Blitum capitatum'' – strawberry blite, blite goosefoot, strawberry goosefoot, strawberry spinach, Indian paint, Indian ink ** '' Blitum virgatum'' (Syn. ''Chenopodium foliosum'') – leafy goosefoot * '' Chenopodiastrum'' (5 species): ** '' Chenopodiastrum murale'' – nettle-leaved goosefoot ** '' Chenopodiastrum simplex'' – giant seed goosefoot * '' Dysphania'' (about 43 glandular species, as ''C. botrys, C. carinatum, C. cristatum, C. melanocarpum, C. multifidium, C. pumilio'' and more) * '' Lipandra'' (one species): ** ''Lipandra polysperma'' – many-seeded goosefoot * '' Oxybasis'' (5 species): ** '' Oxybasis chenopodioides'' – small red goosefoot, saltmarsh goosefoot ** '' Oxybasis glauca'' – oak-leaved goosefoot ** '' Oxybasis rubra'' – red goosefoot, coastblite goosefoot ** '' Oxybasis urbica'' – upright goosefoot * '' Teloxys'' (one species): ** ''Teloxys aristata'' * '' Suaeda australis'' – austral seablite (as ''C. australe, C. insulare'')

Ecology

Certain species grow in large thickets, providing cover for small animals. Goosefoot foliage is used as food by the

caterpillar Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths). As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder ...

s of certain

Lepidoptera Lepidoptera ( ) or lepidopterans is an order (biology), order of winged insects which includes butterflies and moths. About 180,000 species of the Lepidoptera have been described, representing 10% of the total described species of living organ ...

. The seeds are eaten by many

bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...

s, such as the

yellowhammer The yellowhammer (''Emberiza citrinella'') is a passerine bird in the Emberiza, bunting family that is native to Palearctic, Eurasia and has been introduced species, introduced to New Zealand and Australia. Most European birds remain in the br ...

(''Emberiza citrinella'') of Europe or the white-winged fairy-wren (''Malurus leucopterus'') of

Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...

. Goosefoot

pathogens In biology, a pathogen (, "suffering", "passion" and , "producer of"), in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a germ. The term ...

include the positive-sense ssRNA viruses apple stem grooving virus, sowbane mosaic virus and tobacco necrosis virus.

Uses

The genus ''Chenopodium'' contains several plants of minor to moderate importance as food crops as

leaf vegetable Leaf vegetables, also called leafy greens, vegetable greens, or simply greens, are plant leaves eaten as a vegetable, sometimes accompanied by their petioles and shoots, if tender. Leaf vegetables eaten raw in a salad can be called salad gre ...

s – used like the closely related

spinach Spinach (''Spinacia oleracea'') is a leafy green flowering plant native to Central Asia, Central and Western Asia. It is of the order Caryophyllales, family Amaranthaceae, subfamily Chenopodioideae. Its leaves are a common vegetable consumed eit ...

(''Spinacia oleracea'') and similar plants called ''quelite'' in

Mexico Mexico, officially the United Mexican States, is a country in North America. It is the northernmost country in Latin America, and borders the United States to the north, and Guatemala and Belize to the southeast; while having maritime boundar ...

and

pseudocereal A pseudocereal or pseudograin is one of any non-grasses that are used in much the same way as cereals (true cereals are grasses). Pseudocereals can be further distinguished from other non-cereal staple crops (such as potatoes) by their being pr ...

s. These include white goosefoot (''C. album''), '' kañiwa'' (''C. pallidicaule'') and

quinoa Quinoa (''Chenopodium quinoa''; , from Quechuan languages, Quechua ' or ') is a flowering plant in the Amaranthaceae, amaranth family. It is a herbaceous annual plant grown as a crop primarily for its edible seeds; the seeds are high in prote ...

(''C. quinoa''). On the

island of

Crete Crete ( ; , Modern Greek, Modern: , Ancient Greek, Ancient: ) is the largest and most populous of the Greek islands, the List of islands by area, 88th largest island in the world and the List of islands in the Mediterranean#By area, fifth la ...

, tender shoots and leaves of a species called ''krouvida'' () or ''psarovlito'' () are eaten by the locals, boiled or steamed. As studied by Bruce D. Smith, Kristen Gremillion and others, goosefoots have a history of culinary use dating back to 4000 BC or earlier, when pitseed goosefoot (''C. berlandieri'') was a staple crop in the Native American Eastern Agricultural Complex, and when white goosefoot was apparently used by the Ertebølle culture of

Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...

. Members of the eastern European Yamnaya culture also harvested white goosefoot as an apparent cereal substitute to round out an otherwise mostly meat and dairy diet c.'' ''3500–2500'' ''BC. There is increased interest in particular in goosefoot seeds today, which are suitable as part of a

gluten-free diet A gluten-free diet (GFD) is a nutritional plan that strictly excludes gluten, which is a mixture of prolamin proteins found in wheat (and all of its species and hybrids, such as spelt, kamut, and triticale), as well as barley, rye, and oats ...

. Quinoa oil, extracted from the seeds of ''C. quinoa'', has similar properties, but is superior in quality, to corn oil. Oil of chenopodium is extracted from the seeds of

, which is not in this genus anymore.

Shagreen Shagreen is a type of rawhide consisting of rough untanned skin, historically from a horse's or onager's back, or from shark or ray. Etymology The word derives from the French ' and is related to Italian and Venetian ''sagrin'', deriv ...

leather was produced in the past using the small, hard goosefoot seeds. ''C. album'' was one of the main

model organism A model organism is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the model organism will provide insight into the workings of other organisms. Mo ...

s for the molecular biological study of chlorophyllase. Goosefoot

pollen Pollen is a powdery substance produced by most types of flowers of seed plants for the purpose of sexual reproduction. It consists of pollen grains (highly reduced Gametophyte#Heterospory, microgametophytes), which produce male gametes (sperm ...

, in particular of the widespread and usually abundant ''C. album'', is an

allergen An allergen is an otherwise harmless substance that triggers an allergic reaction in sensitive individuals by stimulating an immune response. In technical terms, an allergen is an antigen that is capable of stimulating a type-I hypersensitivi ...

to many people and a common cause of

hay fever Allergic rhinitis, of which the seasonal type is called hay fever, is a type of rhinitis, inflammation in the nose that occurs when the immune system overreacts to allergens in the air. It is classified as a Allergy, type I hypersensitivity re ...

. The same species, as well as some others, have seeds which are able to persist for years in the soil seed bank. Many goosefoot species are thus significant

weed A weed is a plant considered undesirable in a particular situation, growing where it conflicts with human preferences, needs, or goals.Harlan, J. R., & deWet, J. M. (1965). Some thoughts about weeds. ''Economic botany'', ''19''(1), 16-24. Pla ...

s, and some have become

invasive species An invasive species is an introduced species that harms its new environment. Invasive species adversely affect habitats and bioregions, causing ecological, environmental, and/or economic damage. The term can also be used for native spec ...

. In Australia, the larger ''Chenopodium'' species are among the plants called "bluebushes". According to the 1889 book ''The Useful Native Plants of Australia'', ''Chenopodium auricomum'' "is another of the salt-bushes, which, besides being invaluable food for stock, can be eaten by man. All plants of the Natural Order Chenopodiaceae (Salsolacese) are more or less useful in this respect." The book goes on to give the following account from the ''Journal de la Ferme et des Maisons de campagne'':

We have recently gathered an abundant harvest of leaves from two or three plants growing in our garden. These leaves were put into boiling water to blanch them, and they were then cooked as an ordinary dish of spinach, with this difference in favour of the new plant, that there was no occasion to take away the threads which are so disagreeable in chicory, sorrel, and ordinary spinach. We partook of this dish with relish—the flavour—analogous to spinach, had something in it more refined, less grassy in taste. The cultivation is easy: sow the seed in April (October) in a well-manured bed, for the plant is greedy; water it. The leaves may be gathered from the time the plant attains 50 centimetres (say 20 inches) in height. They grow up again quickly. In less than eight days afterwards another gathering may take place, and so on to the end of the year.

Safety

Sphaeraphides occur in the leaves, stem, pith and mesophloem.

Fossil record

†''Chenopodium wetzleri'' fossil seeds of the Chattian stage,

Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...

, are known from the Oberleichtersbach Formation in the

Rhön Mountains The Rhön Mountains () are a group of low mountains (or ''Mittelgebirge'') in central Germany, located around the border area where the states of Hesse, Bavaria and Thuringia come together. These mountains, which are at the extreme southeast end o ...

, central

Germany Germany, officially the Federal Republic of Germany, is a country in Central Europe. It lies between the Baltic Sea and the North Sea to the north and the Alps to the south. Its sixteen States of Germany, constituent states have a total popu ...

.The floral change in the tertiary of the Rhön mountains (Germany) by Dieter Hans Mai - Acta Paleobotanica 47(1): 135-143, 2007.

References

Susy Fuentes-Bazan, Pertti Uotila, Thomas Borsch: ''A novel phylogeny-based generic classification for Chenopodium sensu lato, and a tribal rearrangement of Chenopodioideae (Chenopodiaceae).'' In: ''Willdenowia.'' Vol. 42, No. 1, 2012, p. 5-24. Gelin Zhu, Sergei L. Mosyakin & Steven E. Clemants
''Chenopodium'' - In: Wu Zhengyi, Peter H. Raven, Deyuan Hong (Hrsg.): Flora of China.
Volume 5: Ulmaceae through Basellaceae. Science Press/Missouri Botanical Garden Press, Beijing/St. Louis 2003, , p. 378-.